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The Arctic Warbler Phylloscopus borealis– three anciently separated cryptic species revealed
Authors:PER ALSTRÖM  ISAO NISHIUMI  YOSHIMITSU SHIGETA  KEISUKE UEDA  MARTIN IRESTEDT  MATS BJÖRKLUND  URBAN OLSSON
Institution:1. Swedish Species Information Centre, Swedish University of Agricultural Sciences, Box 7007, SE‐750 07 Uppsala, Sweden;2. Department of Zoology, National Museum of Nature and Science, 3‐23‐1 Hyakunin‐cho, Shinjuku‐ku, Tokyo 169‐0073, Japan;3. Yamashina Institute for Ornithology, 115 Konoyama, Abiko, Chiba 270‐1145, Japan;4. Department of Life Science, Rikkyo University, 3‐34‐1 Nishi‐ikebukuro, Toshima‐ku, Tokyo 171‐8501, Japan;5. Molecular Systematics Laboratory, Swedish Museum of Natural History, PO Box 50007, SE‐104 05 Stockholm, Sweden;6. Department of Animal Ecology, Uppsala University, Norbyv?gen 18D, SE‐752 36 Uppsala, Sweden;7. Department of Zoology, University of G?teborg, Box 463, SE‐405 30 G?teborg, Sweden
Abstract:The Arctic Warbler Phylloscopus borealis breeds across the northern Palaearctic and northwestern‐most Nearctic, from northern Scandinavia to Alaska, extending south to southern Japan, and winters in Southeast Asia, the Philippines and Indonesia. Several subspecies have been described based on subtle morphological characteristics, although the taxonomy varies considerably among different authors. A recent study (T. Saitoh et al. (2010) BMC Evol. Biol. 10 : 35) identified three main mitochondrial DNA clades, corresponding to: (1) continental Eurasia and Alaska, (2) south Kamchatka, Sakhalin and northeast Hokkaido, and (3) most of Japan (Honshu, Shikoku, Kyushu). These three clades were estimated to have diverged during the late Pliocene to early Pleistocene (border at c. 2.6 million years ago). Differences in morphometrics have also been reported among members of the three clades (T. Saitoh et al. (2008) Ornithol. Sci. 7 : 135–142). Here we analyse songs and calls from throughout the range of the Arctic Warbler, and conclude that these differ markedly and consistently among the populations representing the three mitochondrial clades. Kurile populations, for which no sequence data are available, are shown to belong to the second clade. To determine the correct application of available scientific names, mitochondrial DNA was sequenced from three name‐bearing type specimens collected on migration or in the winter quarters. Based on the congruent variation in mitochondrial DNA, morphology and vocalizations, we propose that three species be recognized: Arctic Warbler Phylloscopus borealis (sensu stricto) (continental Eurasia and Alaska), Kamchatka Leaf Warbler Phylloscopus examinandus (Kamchatka (at least the southern part), Sakhalin, Hokkaido and Kurile Islands), and Japanese Leaf Warbler Phylloscopus xanthodryas (Japan except Hokkaido).
Keywords:cryptic species  cytochrome‐b  phylogeny  taxonomy  vocalizations
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